Theor Ecol (2008) 1:2128

DOI 10.1007/s12080-007-0002-0

ORIGINAL PAPER

A general theory of ecology

Samuel M. Scheiner & Michael R. Willig

Received: 9 January 2007 / Accepted: 22 May 2007 / Published online: 29 June 2007
# Springer Science + Business Media B.V. 2007

Abstract Ecologists bemoan the dearth of theory in ecology, in particular, the lack of an overarching, general theory.
These complaints largely are unjustified. The components of
a general theory of ecology have existed for the past half
century; ecologists simply have failed to explicitly recognize
them. We present a general theory of ecology and show how
it relates to ecologys numerous constituent theories and
models. The general theory consists of a description of
the domain of ecology and a set of fundamental principles.
The domain of ecology is the spatial and temporal patterns of
the distribution and abundance of organisms, including
causes and consequences. Fundamental principles are broad
statements about the patterns that exist and the processes that
operate within a domain. The seven fundamental principles
of the theory of ecology are: the heterogeneous distribution
of organisms, interactions of organisms, contingency, environmental heterogeneity, finite and heterogeneous resources,
the mortality of organisms, and the evolutionary cause of
ecological properties. These principles are the necessary and
sufficient elements for a general theory of ecology. The
propositions of any constituent theory of ecology can be
shown to be a consequence of these fundamental principles
along with principles from other science domains. The
S. M. Scheiner (*)
Division of Environmental Biology,
National Science Foundation,
4201 Wilson Blvd.,
Arlington, VA 22230, USA
e-mail: sscheine@nsf.gov
M. R. Willig
Center for Environmental Sciences and Engineering
and Department of Ecology and Evolutionary Biology,
University of Connecticut,
Storrs, CT 06269, USA
e-mail: michael.willig@uconn.edu

general theory establishes relationships among constituent

theories through shared fundamental principles. The next
challenge is to develop and integrate unified, constituent
theories and to establish the relationships among them within
the framework established by the general theory.
Keywords Conceptual framework . law . model . theory

Introduction
Like the emperor who had no clothes, ecology often is
criticized as lacking the theoretical adornments of the
physical sciences. Indeed, for many years ecologists have
decried the fact that we have no general theories. Often, this
debate is framed as whether ecology has laws (e.g., Lawton
1999; Simberloff 2004), a necessary component of a welldeveloped theory. We contend that pessimism about the
theoretical foundations of ecology is ill founded: ecology
has had a robust theoretical framework for many years. We
ecologists simply have not recognized that fact, in part
because we have misunderstood the nature and form of a
comprehensive theory. Given a proper understanding of the
nature and form of an overarching theory, exposing a
general theory of ecology becomes less challenging. The
goal of this paper is to present the fundamental principles of
such a theory so as to further its development.

What is a theory?
A theory is a framework or system of concepts and
propositions that provides causal explanations of phenomena within a particular domain (Hempel 1965; Suppe 1977;
Miller 1987; Giere 1988). The purpose of a theory is to

22

provide a set of linkages for observations and lower-level

models or theories of those observations. Confusion exists
because the term theory is used in reference to a variety
of kinds of systems at different levels of specificity.
Moreover, the nature of theory differs at each level. In this
context, we recognize three levels of theory.
At the broadest level, a general theory consists of an
entire domain of science and a set of fundamental principles
(e.g., the theory of evolution, Darwin 1859; Mayr 1982;
Kutschera and Niklas 2004). That is the type of theory that
we present, a general theory of ecology. A general theory
does not make specific predictions. Rather, it provides the
scaffolding on which the components of more specific,
constituent theories are assembled and integrated. It
exposes assumptions that are sometimes hidden at the level
of models or constituent theories and identifies areas that
are ripe for theoretical development. In addition, it provides
the interconnections that link constituent theories to each
other. A mature or fully developed general theory enhances
understanding at the broadest of levels and facilitates
prediction and forecasting via well articulated models and
constituent theories.
At the intermediate level are constituent theories, which
set boundaries and identify parameters for particular
interest, guiding model development. Constituent theories
can overlap in domain and differ in scope. Depending on
the form and domain of a constituent theory, it may make
no predictions, or it may make qualitative predictions.
Most important, a constituent theory unifies a set of
interrelated models. For example, Scheiner and Willig
(2005) present a constituent theory in ecology that pertains
to variation in species richness along environmental
gradients and show how a number of models are derived
from its propositions.
At the most specific level, models are instantiations of a
constituent theory. At this level, predictions are formalized
and causal understanding motivates the process. Most
references to theory in ecology actually relate to one of
these instantiations (Ives and Agrawal 2005, and related
papers). We prefer the word model for these theories to
indicate that the domain is relatively narrow and predictions
are contingent on well-specified conditions. Models often
are referred to as theories and are the theories that are most
familiar to ecologists. We emphasize that this is not the
level of theory that we present here.
If quantitative predictions occur at the level of models,
what is the utility of the other levels? That question is easy
to answer for constituent theories. Such theories identify the
necessary structure for models to address a particular issue
(e.g., diversity along gradients). Such guidelines make
model development easier and more complete, and can
reveal interrelationships among seemingly disparate models. For example, Scheiner and Willig (2005) demonstrate

Theor Ecol (2008) 1:2128

how an articulated theory can be used to guide model

development.
The issue of prediction is separate from the existence
of a well-developed theory. A theory can provide
understanding without prediction. Precise predictions
may be difficult or impossible for many reasons, including
context dependency, scale dependency, nonequilibrial
dynamics, and the importance of historical contingency.
Ecological theoriesboth general and constituentare
often probabilistic rather than deterministic, identifying
the contexts within which particular patterns or process
are more likely to manifest. To the extent that theory
drives research, constituent theories make clear where
assumptions lie, the sorts of predictions that are possible,
and criteria for evaluating the relative merits of different
lines of inquiry.
The utility of a general theory may be less apparent
because it is two levels removed from the quantitative
predictions that rigorously evaluate theories and test models
(Ives and Agrawal 2005). Consequently, we address this
issue at the end of this essay, once we have clarified the
nature of a general theory and have articulated a theory
of ecology.

A general theory of ecology

A general theory consists of a domain plus a set of
fundamental principles. Fundamental principles are broad
statements about empirical patterns and the processes that
operate within a domain. The domain and fundamental
principles of a general theory of ecology are presented
below. We do not present the fully amplified theory with all
of its components. Such an endeavor would take an entire
book. Rather, our goal is to present its basic premises in
sufficient detail so that others understand their meaning and
reasons for inclusion within the theory. Clarification of the
nature of those premises results in a framework for theories
in ecology. Having such a framework and articulating the
theories within it will help to catalyze the unification of
theory in ecology.
We do not claim that the general theory of ecology is
novel. Quite the contrary, its elements have existed for at
least the past 50 years. However, it has never been
described formally and concisely, although many of its
principles are implicit in the table of contents of most
ecology textbooks (see references in Table 1). Nor do we
claim that the theory, as presented here, is the final version.
Rather, it should be considered provisional and evolving.
The list of fundamental principles may require additions,
deletions, or refinements. Critically, this process can occur
only after the theory has been formalized and discussed in
an open scientific forum.

Theor Ecol (2008) 1:2128

Table 1 Definitions of ecology as presented in various textbooks
Source

Definition

Odum (1971)

[T]he study of the structure and function

of nature
[The] scientific study of the relationships between
organisms and their environments

McNaughton
and Wolf
(1973)
Ricklefs (1979)

Colinvaux
(1986)
Ehrlich and
Roughgarden
(1987)
Stiling (1992)
Dodson et al.
(1998)
Krebs (2001)

Begon et al.
(2006)
Gurevitch et al.
(2006)

This paper

[T]he study of the natural environment,

particularly the interrelationships between
organisms and their surroundings
[T]he study of animals and plants in relation to
their habits and habitats
[T]he study of the relationship between organisms
and their physical and biological environments
[T]he study of interactions between organisms and
between organisms and their environments
[T]he study of the relationships, distribution, and
abundance of organisms, or groups of organisms,
in an environment
[T]he scientific study of the interactions that
determine the distribution and abundance of
organisms
[T]he scientific study of the interactions between
organisms and their environment
[T]he study of the relationships between living
organisms and their environments, the
interactions of organisms with one another, and
the patterns and causes of the abundance and
distribution of organisms in nature
The study of the spatial and temporal patterns of
the distribution and abundance of organisms,
including causes and consequences

The domain of ecology

Before we synthesize and make explicit the structure of
this general theory, we need to define its domain.
Although domains are artificial constructs, they serve as
foci for organizing theories into coherent entities. The
domain of ecology and its general theory is the spatial
and temporal patterns of the distribution and abundance
of organisms, including causes and consequences.
Although our definition of the domain spans the
definitions found in most textbooks (Table 1), it is
different in two respects. First, our definition includes the
phenomena to be explained (i.e., spatial and temporal
patterns of abundance of organisms) and the causes of
those phenomena. Some definitions include only the latter
(i.e., interactions of organisms and environments). Second,
and most strikingly, our definition explicitly includes the
study of the consequences of those phenomena, thereby
embracing much of ecosystem sciences.

23

The rules governing fundamental principles

Fundamental principles can be determined by a simple set
of rules, although we recognize that the application of the
rules is open to debate and disagreement. Here, we present
the rules in the abstract; they are made more concrete when
applied to the principles of the theory of ecology.
Fundamental principles are of two general types: those
unique to the domain of consideration, and those shared
with other domains. If a fundamental principle is unique to
a domain, it must meet one of two criteria, either the
principle is shared by many constituent theories within the
domain, or the principle is necessary for distinguishing
competing general theories. We refer to the former criterion
as the inclusionary rule and the latter as the exclusionary
rule. Inclusionary principles must be broad, whereas
exclusionary principles may be narrow and can often be
the result of the history of debates about a theory. If a
fundamental principle is shared with another domain, it
must be a consequence of mechanisms from another
domain and have domain-specific causal significance. We
refer to this criterion as the causal rule.
These criteria mean that a principle taken unchanged
from another domain need not be specified within a theory.
We take as given the fundamental principles of any other
general theory. We recognize the general tenet of consilience, that the entire set of scientific theories must be
consistent with each other (Whewell 1858). Theories may
clash, but such clashes indicate foci of research that
advance understanding. In general, theories inhabiting
different domains will not clash directly, although results
from one domain can point to problems with theories in
other domains. For example, studies of geographical
distributions of clades of organisms, within the domain of
historical biogeography, became important evidence for the
theory of continental drift, a part of the domain of geology.
In that instance, the need for a causal mechanism to explain
distribution patterns (the causal rule) was one factor that led
to the development of new fundamental principles in
another domain.
The fundamental principles of ecology
The general theory of ecology consists of seven fundamental principles (Table 2). These principles were in place by
the 1950s and were widely accepted by most ecologists
after the coalescence of the field of ecosystem ecology and
the cementing of the Modern Synthesis in evolutionary
biology. Of course, the roots of all of the principles go back
much further.
The first fundamental principlethe heterogeneous
distribution of organismsis a refinement of the domain.
It encompasses the basic object of interest and its most

24
Table 2 The seven fundamental principles of the general theory of
ecology and the rules that determine their inclusion
Seven fundamental principles
1. Organisms are distributed in space and time in a heterogeneous
manner. [inclusionary rule]
2. Organisms interact with their abiotic and biotic environments.
[inclusionary rule]
3. The distributions of organisms and their interactions depend on
contingencies. [exclusionary rule]
4. Environmental conditions are heterogeneous in space and time.
[causal rule]
5. Resources are finite and heterogeneous in space and time.
[causal rule]
6. All organisms are mortal. [causal rule]
7. The ecological properties of species are the result of evolution.
[causal rule]

important property. The heterogeneity of distribution is one

of the most striking features of nature: all species have a
heterogeneous distribution at some, if not most, spatial
scales. Arguably, the origins of ecology as a discipline and
the first ecological theories can be traced to its recognition
(Forster 1778; von Humboldt 1808). This heterogeneous
distribution is both caused by and a cause of other
ecological processes. The fundamental principles are not
independent causal mechanisms, rather, the mechanisms
that they encompass interact.
The second fundamental principleinteractions of
organismsincludes within it the vast majority of ecological processes responsible for heterogeneity in time and
space. Many definitions of ecology are equivalent to this
principle (Table 1). Within this principle, particular interactions that are part of constituent theories act to unpack the
general theory (see Constituent theories section).
The third fundamental principlecontingencyrepresents
either the inclusionary rule or the exclusionary rule, depending
on views of the history of theories in ecology. We discuss the
exclusionary nature of this principle at the time of its genesis
(see Competing general theories section). Since that time,
recognition of the importance of contingency in all ecological
processes has increased steadily and now appears in a wide
variety of constituent theories and models. Contingency is an
important cause of the heterogeneous distribution of
organisms, both at very large extents of time and space
(e.g., a particular species arose on a particular continent) and
at very small extents (e.g., a seed lands in one spot and not
another).
We speculate that general theories of all domains include
some version of this principle (e.g., genetic drift in
evolution; quantum theory in physics). Thus, it might be
considered fundamental to all scientific theories. This
principle demonstrates that the decision to include a

Theor Ecol (2008) 1:2128

fundamental principle and the type of rule that is the basis

for inclusion are not always clear and may be subject to
debate by scientists, historians, and philosophers.
The fourth fundamental principleenvironmental
heterogeneityis a consequence of processes from the
domains of the earth and space sciences. For example,
seasonal variation in temperature is the result of orbital
properties of the Earth, whereas a variety of geophysical
processes create heterogeneity in environmental stressors
like salt (e.g., wave action near shores) or heavy metals
(e.g., geologic processes that create differences in bedrocks). It is beyond the scope of this paper to detail all of
those processes and their domains. Indeed, this principle
encompasses many constituent theories and contains a
broad class of underlying mechanisms for the heterogeneous distribution of organisms. As with the second
principle, particular mechanisms pertain to particular
constituent theories. Again, the fundamental principle
captures a wide range of theories and mechanisms so as
to provide a unifying framework.
The fifth principlefinite and heterogeneous resources
is again a consequence of processes from the domains of the
earth and space sciences. Although variation in resources is
similar to variation in environmental conditions, a fundamental distinction is the finite nature of these resources.
Unlike an environmental condition, a resource is subject to
competition. For example, seasonal variation in light and
temperature are caused by the same orbital mechanisms, but
light is subject to competition (e.g., one plant shades
another), whereas temperature is a condition and not subject
to competition. This distinction in the nature of environmental factors with regard to competitive processes can
result in very different ecological outcomes (e.g., patterns of
diversity in plant communities, Scheiner and Rey-Benayas
1994). Whether a particular environmental factor is a
condition or a resource can be context dependent. For
example, water is sometimes a resource subject to competition (e.g., plants in a desert), and sometimes a condition
(e.g., fish in the ocean).
The sixth fundamental principlethe mortality of
organismsis the result of processes that come from the
domain of organismal biology, physiology, and development. By mortal we mean that no organism is invulnerable, i.e., any organism might die as the result of predation,
stress, or trauma. We do not mean by this principle that all
organisms senesce. The senescence of organisms is a more
narrow version of this principle that would apply to
particular constituent theories. Although the majority of
multicellular species apparently senesce, this has not been
demonstrated for some multicellular species. We are not
aware of an articulated theory of development or physiology that predicts the necessity of or conditions for
senescence, but suspect that it could be accomplished given

Theor Ecol (2008) 1:2128

current knowledge of organisms. We will let philosophers

argue whether a bacterium that splits into two represents a
single, immortal organism or (our position) the end of one
individual and the creation of two new individuals. This
fifth principle, either in the more general version of
vulnerability or in the more narrow version of senescence,
forms the basis of a large number of constituent theories
concerning phenomena as wide ranging as life histories,
behavior, demography, and succession.
The seventh principlethe evolutionary cause of ecological propertiesis the result of processes that derive
from the theory of evolution. The inclusion of evolution
within ecological thinking was an important outcome of the
Modern Synthesis. Although evolutionary thinking about
ecological processes goes back at least to Darwin (1859),
evolutionary thinking had been infusing ecology more
widely at least since the 1920s (Collins 1986; Mitman
1992) and its widespread acceptance occurred primarily in
the latter half of the 20th century. The acceptance of this
principle led to such disciplines as behavioral ecology and
population biology, and the demise of the Clementsian
superorganism theory (see Competing constituent theories).
It is in this latter capacity that this principle can be
considered under the exclusionary rule and the causal rule.
The scope of the fundamental principles
These fundamental principles are necessary and sufficient
to support a general theory of ecology. One or more of the
propositions of each constituent theory of ecology can be
shown to be a consequence either of the fundamental
principles of ecology, or of principles from other domains.
A proposition of a constituent theory can be a consequence
of a fundamental principle in three ways. First, a proposition may be a member of a class defined by a fundamental
principle. For example, competition, a type of biotic
interaction (principle 2), plays a key role in plant
succession (Pickett et al. 1987). Second, a constituent
theory can identify, define, or refine particular patterns or
mechanisms identified in a fundamental principle. For
example, variation in dispersal distance, a form of contingency (principle 3), is a component of population dynamic
theory (Turchin 2001). Third, a proposition in a constituent
theory might be derived formally from a fundamental
principle. For example, in food web theory, rules governing
food chain length derive from aspects of the second law of
thermodynamics (a principle in the domain of physics). In
the next section, we examine further the relationship
between fundamental principles and the propositions of
constituent theories.
To the extent that the fields of ecology can be
characterized as a collection of related or overlapping
constituent theories, these principles provide the scaffolding

25

for a mature and fully developed theory that applies to all

subdisciplines of ecology. The general theory establishes
relationships among constituent theories. Linkages occur
when constituent theories are derived from one or more of
the same fundamental principles. Constituent theories that
share fundamental principles must be consistent with each
other. If they are not, one or more of those theories may
require revision. Some of the propositions may be the
consequence of theories in other domains. To be a
constituent theory within the domain of ecology, at least
one of its propositions would have to be a consequence of
the fundamental principles of the general theory of ecology.
The fundamental principles do not include ecological
entities or systems, such as populations, communities, or
ecosystems. Rather, those entities are a consequence of the
mechanisms embodied by the principles. For example, a
population and its properties are the consequence of
organisms of a single species inhabiting a particular
location (principle 1) and interacting with each other
(principle 2). Philosophers refer to this derivation of one
object from other principles as ontological reduction.
Ontological reduction is favored because it results in theory
simplification and eliminates ontological primitives that
require special explanation.
The fundamental principles do not specify mechanisms.
It is the function of constituent theories to explicate the
types and appropriateness of particular mechanisms. For
example, interactions such as predation, herbivory, competition, mutualism or parasitism might be mechanisms within
theories of community structure. Such lack of specificity
has two causes. The first relates to the diversity of
environmental and historical circumstances which determine the importance of any suite of mechanisms effecting
the properties of ecological systems. Any attempt to
enumerate the circumstances in which particular biotic
interactions dominate would create an unwieldy theory. The
second cause relates to the history of ecology, which can be
characterized as the accumulation of mechanisms about, and
an appreciation for, the complexity of ecological systems.
Rather than debates about the existence of mechanisms,
disagreements mostly have been about their relative importance, a dispute that occurs at the level of constituent
theories, rather than at the level of the general theory.
The exact list of fundamental principles of ecology
certainly will be the subject of debate. Recently, a partial
form of our general theory was advanced by Lawton (1999)
as a list of universal laws that underpin all ecological
systems that are equivalent to principles 2 and 7. He
included other principles such as the first and second laws
of thermodynamics and the conservation of mass. We did
not include those principles because they were taken
unchanged from the domain of physics. He also included
physical principles governing a broad set of properties

26

which underpin the physiology of organisms. Those were

not included because we consider the physiology of
organisms to be outside the domain of ecology. The effects
of principles in that domain appear in our theory in the
guise of principle 6. Otherwise, those principles might play
roles in more specific, constituent theories. Our effort also
differs from that of Lawton by providing a set of rules by
which one can decide whether a principle should be
included within the general theory.

Constituent theories
A general theory of ecology should be able to construct and
accommodate all constituent theories within the domain of
ecology. We demonstrate this accommodation and the way
in which their propositions are consequences of fundamental principles by examining two constituent theories.
Turchin (2001) presented a theory of population dynamics
consisting of various models of population growth, which
he referred to as laws and enunciated several propositions
that form the bases of those laws. The propositions, which
he called foundational concepts, are: exponential population growth, self-limitation, and trophic oscillations. These
propositions are a consequence of fundamental principles
26. The pattern of exponential population growth is a
function of the birth and death (principle 6) of individuals,
but can also be affected by environmental heterogeneity
(principle 4) and various types of contingency (principle 3).
Population self-limitation occurs because resources are
finite (principle 5). Trophic oscillations occur because of
interactions among species (principle 2). Although not
expressed precisely within the conceptual framework
presented here, Turchins ideas amount to a constituent
theory that it is quite mature.
This example also demonstrates how fundamental
principles can set boundaries on the form of constituent
theories and models. Turchin (2001) explains how a system
that contained an additional proposition involving spontaneous generation would result in different models of
population growth, linear rather than exponential. Clearly,
however, the proposition of spontaneous generation is not
supported by any fundamental principle in any general
theory of biology.
Scheiner and Willig (2005) presented a theory of
diversity gradients consisting of four propositions: limiting
resources (a consequence of principle 5), species pools
(principle 1), environmental variation (principle 4), and
population or organismal trade-offs (principles 6 and 7).
Those propositions, in turn, embody a large variety of
models. While those models have been presented as
contradictory or in opposition, when presented as part of a
framework of propositions their commonalities become

Theor Ecol (2008) 1:2128

clear. Levins (1968) pointed out that no model can

simultaneously maximize generality, precision, and realism.
At most, a model can strive to maximize two of these
qualities, and the majority of models are a compromise
among all three. Since there is no best all-purpose model,
theory unification permits one to choose which among a
variety of models is most appropriate for a given situation,
while allowing one to relate model results to those of other
models. In a similar fashion, a general theory exposes
commonalities among constituent theories.
Ecology contains a plethora of constituent theories that
differ in maturity (Table 3). Some, such as the equilibrium
theory of island biogeography, are mature. Substantial
effort has refined the mathematical models of that theory
and demonstrated its empirical utility. Other constituent
theories are promising but underdeveloped. For example,
the metabolic theory of ecology currently consists of a
single model and lacks a broader unifying set of propositions that would facilitate the building of alternative
models.
For any of these constituent theories, we could go
through a similar exercise of explicating their propositions
and showing how those propositions are a consequence of
the fundamental principles of ecology. Our claim is that one
or more of the fundamental principles are necessary to any
of ecologys constituent theories, and the fundamental
principles of the entire set of general theories in science
are sufficient to derive their propositions. It is beyond the
Table 3 Selected constituent theories of ecology
Theory

Stage of
maturity

Reference

Island biogeography theory

More
complete
More
complete
More
complete
More
complete
More
complete
More
complete
Intermediate
Intermediate

MacArthur and
Wilson (1967)
Stephens and
Krebs (1986)
Pickett et al. (1987)

Foraging theory
Succession theory
Metapopulation theory
Population dynamic theory
Niche theory
Food web theory
Theory of latitudinal
gradients of diversity
Metabolic theory of ecology
Unified theory of diversity
gradients
Metacommunity theory

Intermediate
Intermediate
Less
complete

Hanski (1999)
Turchin (2001)
Chase and
Leibold (2003)
Pimm (1982)
Willig et al. (2003)
Brown et al. (2004)
Scheiner and
Willig (2005)
Leibold et al. (2004)

The associated reference is illustrative rather than definitive or

comprehensive.

Theor Ecol (2008) 1:2128

scope of this paper to verify this claim in an exhaustive

fashion, although that is a paramount task for further
development of a general theory of ecology.
Theories differ in their level of generality. We classify a
theory as a constituent theory if it meets one of two criteria:
the theory serves to unite many models that attempt to
explain a single phenomenon (e.g., the theory of diversity
gradients), or a single model is portrayed as explaining a
broad range of phenomena (e.g., the metabolic theory of
ecology). Otherwise, the theory is likely a model within a
more general constituent theory, or a nascent constituent
theory.

The nature and benefits of a general theory of ecology

What we term constituent theory is what ecologists often
refer to as theory with all of its components, including
facts, generalization, laws, models and hypotheses (Pickett
et al. 1994). A general theory is sparser, consisting of a
domain and a set of fundamental principles, which together
define a framework. For a general theory, the other
components are encapsulated within its constituent theories.
A general theory is mature only to the extent that its
constituent theories are mature and their interrelationships
are clear. In this regard, the general theory of ecology is an
adolescent, some of its constituent theories are quite
mature, while others are still under ripe, and the interrelationships among them are not well established.
Competing constituent theories
A general theory provides several benefits. It helps
scientists decide between competing constituent theories.
For example, in plant ecology in the first half of the
twentieth century, two competing theories considered the
nature of communities. These two extreme viewpoints
differed in the importance they ascribed to biotic versus
abiotic factors, and predictable versus random processes in
shaping community structure.
The first theory, proposed by Clements (1916), saw plant
communities as highly organized entities made up of
mutually interdependent species. Communities were superorganismsthe analogue of individual organismsthat are
born, develop, grow, and eventually die. Two of the
hallmarks of the superorganism concept were the presence
of very tight linkages among species within communities
and cooperation among the species in a community for the
benefit of the entire community.
The alternative theory, put forward by Gleason (1917;
1926), posited that communities are the result of the
interactions of each species and the environment (biotic
and abiotic factors) in combination with chance historical

27

events. Each species has its own environmental tolerances

and thus responds in its own way to environmental
conditions. The implication was that along an environmental gradient, different species would have their boundaries
at different places. Not only were communities not tightly
linked superorganisms, but defining the collection of
species living together in a particular place as a community
was an arbitrary human construct.
These two theories disagreed on a number of grounds
that can be seen within the framework of the general theory.
For example, they held to different ideas about how
evolution shapes ecological traits (principle 7). They
differed in the nature of environmental heterogeneity
(principle 4), and they differed in the nature of interactions
among organisms (principle 2). If the general theory of
ecology had been formalized while those theories were
being debated, it might have made the nature of the
disagreements clearer, focused the disagreements along
productive lines of inquiry, and pointed to the evidence
needed to decide between them.

Competing general theories

Formalizing a general theory facilitates an assessment of
the merits of competing general theories. Competing
general theories are distinguished by either requiring,
rejecting, or altering the form of fundamental principles.
The shift in ecology from an equilibrium paradigm to a
nonequilibrium paradigm illustrates this process. In terms
of our general theory, one aspect of this shift was the
recognition of the importance of contingency through the
addition of the third principle. A general theory of ecology
that posits that the world is in equilibrium, while potentially
recognizing a role for contingency, would not see contingency as being a major mechanism. Rather, the equilibrium
paradigm included a strongly deterministic viewpoint, and
principle 3 was simply part of particular models rather than
a fundamental principle. We realize that this example is
oversimplified and does not capture the complexities of the
debate. McIntosh (1985) provides a detailed discussion of
the history of this conceptual shift.
A general theory also highlights the nature of disagreements about mechanisms. In the general theory of ecology,
no biotic mechanism is given primacy. During the 1970s
and 1980s, a debate raged among ecologists about the
primacy of competition in structuring communities (McIntosh
1985). The outcome of that debate was an agreement that
competition is an important process, but that other processes
can be of equal or greater importance in many circumstances.
In the context of the general theory of ecology, the debate
was whether the set of fundamental principles was insufficient and needed an additional statement about the primacy

28

of competition, with the conclusion being that no such

principle was warranted.

Whither ecology?
We recognize that ecology presents challenges for unificationthe vast number of objects (e.g., taxa) to be
considered leads to an extreme diversity of disparate
models. All the more reason to reiterate our previous call
for ecologists to develop and integrate unified, constituent
theories (Scheiner and Willig 2005). The general theory
presented here should help guide that process. We ecologists need no longer argue among ourselves about the
strength of the theoretical underpinnings of our discipline.
The components of a general theory have existed for many
years. The next challenge is to establish the relationships
among ecologys constituent theories. Indeed, the cries that
the empress has no clothes were myopic. Now fully vested,
it is clear: ecology is a well dressed lady, as elegant as any
of her sister disciplines in the sciences.
Acknowledgements For insightful comments, we thank Jim Collins,
Gordon Fox, Jim Grace, Bob Holt, Jane Maienschein, Alan Tessier,
Bill Zamer, and several anonymous reviewers. This manuscript is
based on work done by both authors while serving at the National
Science Foundation. The views expressed in this paper do not
necessarily reflect those of the National Science Foundation or the
United States Government. Partial support to MRW was provided by
the Center for Environmental Sciences and Engineering at the
University of Connecticut.

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